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On Elevation-Related Shifts of Spring Activity in Male Vipers of the Genera ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Herpetozoa Jahr/Year: 2019 Band/Volume: 31_3_4 Autor(en)/Author(s): Stümpel Nikolaus, Zinenko Oleksander, Mebert Konrad Artikel/Article: on elevation-related shifts of spring activity in male vipers of the genera Montivipera and Macrovipera in Turkey and Cyprus 125-132 StuempelZinenkoMebert_Spring_activity_Montivipera-Macrovipera:HERPETOZOA.qxd 12.02.2019 15:04 Seite 1 HERPEToZoA 31 (3/4): 125 - 132 125 Wien, 28. Februar 2019 on elevation-related shifts of spring activity in male vipers of the genera Montivipera and Macrovipera in Turkey and Cyprus (squamata: serpentes: Viperidae) Zur höhenabhängigen Frühjahrsaktivität männlicher Vipern der Gattungen Montivipera und Macrovipera in der Türkei und Zypern (squamata: serpentes: Viperidae) NikolAus sTüMPEl & o lEksANdR ZiNENko & k oNRAd MEbERT kuRZFAssuNG der zeitliche Ablauf von lebenszyklen wechselwarmer Wirbeltiere wird in hohem Maße vom Temperaturregime des lebensraumes bestimmt. in Gebirgen sinkt die umgebungstemperatur mit zunehmender Höhenlage. deshalb liegt es nahe, anzunehmen, daß die Höhenlage den Zeitpunkt des beginns der Frühjahres - aktivität von Vipern beeinflußt. um diesen Zusammenhang zu untersuchen, haben die Autoren im Zeitraum von 2004 bis 2015 in der Türkei und auf Zypern den beginn der Frühjahrshäutung bei männlichen Vipern der Gattun - gen Montivipera und Macrovipera zwischen Meereshöhe und 2300 m ü. M. untersucht. sexuell aktive Männchen durchlaufen nach der Winterruhe und vor der Paarung eine obligatorische Frühjahrshäutung. im Häutungsprozeß werden äußerlich klar differenzierbare stadien durchschritten, von denen die Eintrübung des Auges besonders auffällig und kurzzeitig ist. dieses stadium ist daher prädestiniert, um den nachwinterlichen Aktivitätsbeginn zwischen Populationen unterschiedlicher Höhenlagen miteinander zu verglei - chen. Während der Freilandstudien wurden insgesamt 15 adulte männliche Vipern mit milchig-opaken Augen angetroffen, welche ihren Häutungsstatus anzeigten. die vorliegenden Ergebnisse zeigen einen signifikanten positiven Zusammenhang zwischen dem beginn der Frühjahreshäutung der untersuchten Vipern und der Höhenlage ihres Fundortes. Je höher der lebensraum lag, desto später im Jahr setzte die Frühjahrshäutung der Männchen ein. diese Ergebnisse können helfen, für die unter - suchten Arten die dauer der überwinterungsperiode in Abhängigkeit von der Höhenlage ihres Vorkommens abzu - schätzen. Nach bestem Wissen der Autoren ist die vorliegende studie die erste untersuchung, die einen Zusammenhang zwischen der Phänologie und der Höhenverbreitung bei eurasischen Vipern aufzeigt. AbsTRACT At a local scale, the timing of periodic life-history events of ectothermic vertebrates is strongly influenced by the ambient temperature. in the mountains, temperature decreases with increasing elevation. This is why the onset of spring activity of vipers can be expected to vary regionally between warm lowland and cold montane climates. To analyze the relationship between elevation and onset of spring activity, the authors surveyed habitats for emerg - ing male vipers of the genera Montivipera and Macrovipera between April and May in Turkey and Cyprus. sexually active male vipers shed soon after leaving their hibernacula and before mating. The male spring shedding is obligatory and therefore represents a chronometer for the onset of the vipers’ surface activity. during field trips between 2004 and 2015, fifteen male vipers were found in habitats between sea level and 2,300 m a.s.l. elevation with milky-opaque eyes reflecting their state of pre-ecdysis. The observed significant positive correlation between onset of the vipers’ spring activity and the elevation of the record site represents an ecological cline. The information can help to forecast duration of hibernation and activity of the vipers in the field. To the best knowledge of the authors, this is the first study about spring phenol - ogy of vipers from the eastern Mediterranean. kEY WoRds Reptilia: squamata: serpentes: Viperidae; Macrovipera lebetina , Montivipera bulgardaghica , Montivipera wagneri , Montivipera xanthina , physiology, behavior, phenology, ecdysis, molting, spring shedding, activity, Turkey, Cyprus iNTRoduCTioN The ecosystems of the Near and viper species out of three genera ( Vipera , Middle East harbor the highest diversity of Montivipera and Macrovipera ) are record - true vipers within Eurasia. Currently, 13 ed from the Turkish territory (e.g., NilsoN StuempelZinenkoMebert_Spring_activity_Montivipera-Macrovipera:HERPETOZOA.qxd 12.02.2019 15:04 Seite 2 126 N. s TüMPEl & o. Z iNENko & k. M EbERT et al. 1988; sTüMPEl & J oGER 2009). With correlates with increased feeding, metabol - a few exceptions, the species within the ic rate and temperature ( sEMliTsCH 1979; genera are allopatrically distributed ( JoGER APPlEbY 1980; G ibsoN et al. 1989). it also 1984). Mountain Vipers, Montivipera spp., varies by reproductive state and cycle, and blunt-nosed Vipers, Macrovipera spp ., respectively, hormonal status, gender and were subjects to studies on zoogeography ontogenetic stage (e.g., kubiE et al. 1978; (JoGER 1984; M EbERT et al. 2016), taxono - NilsoN 1980; JACobsoN 2007; l illYWHiTE my ( NilsoN & A NdRéN 1986) and phylo - 2014 ). other influential factors may in - geny ( sTüMPEl & J oGER 2009; s TüMPEl et clude exposure to dry air and the resulting al. 2016). However, comparatively little is risk of increasing cutaneous evaporative known on their ecology (e.g ., N ilsoN et al. water loss ( lillYWHiTE & M AdERsoN 1982; 1999; ETTliNG et al. 2013, 2016), while MAdERsoN 1984; lillYWHiTE 2006), wear phenological data are completely lacking or and damage of the skin, as well as detach - hidden in grey literature. Macrovipera is ment of ectoparasites ( looMis 1951; HEAT- generally associated with steppe-like to WolE 1999). semi-arid habitats situated between sea during ecdysis, histological process - level and 1,700 m a.s.l. and reaches its es, which occur in the epidermis, affect the western range limit in the Eastern Medi - external appearance of the snake gradually. terranean. species of the genus Monti - However, several stages of the process are vipera are found in a broader ecological very prominent and identifiable. The ecdy - and elevational spectrum. For example, in sis process lasts a minimum of 14 days in western and south-western Anatolia, these captivity (the authors’ personal observa - mountain vipers inhabit Mediterranean and tions) and is visually marked by a dull oromediterra nean biomes, whereas in east - appearance of the snake. Eight days prior ern Anatolia these vipers are associated to the actual skin shedding, the eyes turn with boreal and subalpine irano-Turanian visibly opaque (milky). This prominent floras. external change lasts for at least two days in ectothermic animals, climatic fac - under controlled lab conditions (the tors influence life history traits. Many of authors’ personal observations). Ecdysis them, such as seasonal and diurnal activity, culminates in the shedding of the old outer reproductive performance, growth rate and keratinized epithelial layer about six days metabolic rate depend on the thermal pro - after the eyes have become clear again (see files of the habitats (e.g., duVAll et al. MAdERsoN 1965). The entire process of 1985; VAN WYk 1995). Therefore, temper - skin renewal is influenced by environmen - ature is a relevant environmental factor to a tal factors (temperature, solar radiation) reptile, defining part of its living space and and, e.g., can last about 40 days in Vipera shaping its abiotic components ( WAlkER et berus (l iNNAEus , 1798) at cool northern al. 1994; köRNER 1998; GREENlANd & latitudes (see olssoN et al. 1997), and like - los lEbEN 2001). Along with the increase ly is shorter in warmer regions or climate of elevation, also climatic, spatial, biotic zones. Furthermore, recent evidence from and evolutionary factors change ( MCCAiN the laboratory and field show that ecdysis & GRYTNEs 2010). The most obvious is the can be a synchronous process within snake generally linear decrease in temperature by populations, likely in correlation with envi - an average of approximately 0.68 °C per ronmental factors and/or pheromones, 100 m increase in elevation ( bARRY 2008). which influence physiology including Consequently, the seasonal and temporal reproductive cycles ( lilliWHiTE & s HEEHY activity of vipers inhabiting different eleva - iii 2016 and references therein). intra - tions can be expected to be affected by population synchrony of the first ecdysis extrinsic abiotic factors. following emergence from hibernacula was one such temporal activity is the observed in different snake groups in North shedding process in snakes (ecdysis). in America (e.g., kubiE et al. 1978; FoRd general, ecdysis happens continuously 1996; P ARkER & ANdERsoN 2007) and throughout the lifetime and is associated places this synchronization into a seasonal with growth, whereby a higher frequency context. The pheromones released into the StuempelZinenkoMebert_Spring_activity_Montivipera-Macrovipera:HERPETOZOA.qxd 12.02.2019 15:04 Seite 3 on elevation-related shifts of spring activity in Montivipera and Macrovipera 127 environment during ecdysis inform other across a broad region, if the same stage of snakes in the area about the reproductive ecdysis is compared. Hence, milky eyes status of the newly
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